Hypsilophodont

Hypsilophodontidae (or Hypsilophodontia) is a traditionally used family of ornithopod dinosaurs, generally considerd invalid today. It historically included taxa from across the world, and spanning from the Middle Jurassic until the Late Cretaceous. This inclusive status was supported by some phylogenetic analyses from the 1990s and mid 2000s,[2][3] although there have also been many finding that the family is an unnatural grouping which should only include the type genus, Hypsilophodon, with the other genera being within clades like Thescelosauridae.[4][5][6][7][8][9][10][11][12][13][14] A 2014 analysis by Norman recovered a grouping of Hypsilophodon, Rhabdodontidae and Tenontosaurus, which he referred to as Hypsilophodontia.[15] All other analyses from around the same time have instead found these latter taxa to be within Iguanodontia.[11][16]

Hypsilophodonts
Temporal range: Early Cretaceous, 130–125 Ma
Hypsilophodon Melb Museum email
Hypsilophodon skeleton
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Clade: Clypeodonta
Family: Hypsilophodontidae
Dollo, 1882[1]
Subgroups
Synonyms
  • Hypsilophodontinae Nopsca, 1928
  • Hypsilophodontia Cooper, 1985

Linnaean usage

Marsh laosaurus
Skeleton of Laosaurus consors as drawn by Othniel Marsh (since named Othnielosaurus)

Hypsilophodontidae was named originally in 1882 by Louis Dollo, as a family to include Hypsilophodon and other small ornithopods with a single row of teeth, four pedal digits, and a rhomboid sternum. For several decades after its naming the family only included Hypsilophodon.[3] In 1911 Karl von Zittel published a textbook on vertebrate classifications, in which he included multiple genera in "Hypsilophontidae" (sic for Hypsilophodontidae[17]), including Hypsilophodon, Nanosaurus, Laosaurus and Dryosaurus. Zittel considered the family to unite all taxa that lacked premaxilla teeth, had a single row of maxilla teeth, neck vertebrae which have flat articulations or a flat front and round back, fused sacral vertebrae, a femur shorter than the tibia, 5 fingered manus' and 4 toed peds.[18] Thescelosaurus was named in 1913 by Charles Gilmore, and its skeleton was described in detail by the same author in 1915. Gilmore had originally classified Thescelosaurus within Camptosauridae, but in the 1915 description he determined that it shared far more features with Hypsilophodontidae. He reclassified Laosaurus, Nanosaurus and Dryosaurus in the family Laosauridae, leaving only Thescelosaurus and Hypsilophodon in Hypsilophodontidae. The characteristics of the family were also re-analysed, and Gilmore showed that the premaxilla actually had teeth, a characteristic of the family; the 3rd manus digit had 4 phalanges; the femur was either shorter or longer than the tibia; and dorsal ribs had only a single articulation point.[17]

Thescelosaurus neglectus, CMN
Skeleton of Thescelosaurus edmontonensis in display as preserved

The first expansive analysis on the relationships of Hypsilophodontidae was that of Swinton in 1936, during a redescription of Hypsilophodon from new specimens. The possible hypsilophodonts Geranosaurus and Stenopelix were removed from the clade (then the subfamily Hypsilophodontinae), and considered to be intermediate basal ornithopods, as there were no features linking them to Hypsilophodon. Thescelosaurus was considered within the family, because of the large number of shared features, as well as Dysalotosaurus, from the Kimmeridgian of Tanzania. Laosaurus and Dryosaurus were not considered hypsilophodonts because of their lack of distinguishable features, as Swinton concluded that they were probably in the family Laosauridae, intermediate between Hypsilophodontidae and Iguanodontidae, and were probably synonyms of each other as well.[19] Charles M. Sternberg (1940) considered there to be multiple genera within the family, all sharing fully enamelled teeth, divided into two subfamilies, Hypsilophodontinae and Thescelosaurinae. Within Hypsilophodontinae–grouped by a longer scapula, thinner forelimb and femora shorter than tibiae–Sternberg included Hypsilophodon, Dysalotosaurus, and Parksosaurus (renaming of Thescelosaurus warreni). Only Thescelosaurus was included in Thescelosaurinae, as it had a tibia shorter than the femur.[20]

Peter M. Galton in 1972 re-studied the relationships of taxa within Ornithischia. Thescelosaurus was removed from Hypsilophodontidae because of its short limbs, meaning it was probably not cursorial, unlike all other hypsilophodonts. The presence of premaxilla teeth, once used to diagnose the group, was found to be present in unrelated taxa like Heterodontosaurus, Protoceratops and Silvisaurus. Galton made Hypsilophodontidae paraphyletic, as he considered Thescelosaurus to be a hypsilophodont, but excluded it from the family Hypsilophodontidae. The phylogenetic hypothesis of Galton is shown below. Taxa considered hypsilophodontids are enclosed by green.[21]

Ornithischia

Fabrosaurus -> Echinodon

Heterodontosaurus

Ankylosauria

Stegosauria

Pisanosaurus

Hadrosauridae

Iguanodon -> Tenontosaurus

Ceratopsia

Stenopelix -> Psittacosaurus

Camptosaurus leedsi -> Dysalotosaurus -> Dryosaurus -> Wealden ornithopod

Camptosaurus

Yaverlandia -> Stegoceras -> Pachycephalosaurus

Laosaurus

Parksosaurus

Hypsilophodon

Laosaurus minimus

Thescelosaurus

Cladistic usage

Question of monophyly

in 1992 David Weishampel and Ronald Heinrich reviewed the systematics and phylogenetics of Hypsilophodontidae. Hypsilophodontidae was supported as a monophyletic clade that encompassed "thescelosaurids", Hypsilophodon and Yandusaurus. The family was diagnosed by the absence of ridges that end as denticles in teeth (reversed in Hypsilophodon); presence of a single central ridge on dentary teeth; ossified sternal plates on torso ribs; and a straight and unexpanded shape of the prepubis. Their resulting cladogram is reproduced below:[3]

Euornithopoda
Iguanodontia

Tenontosaurus

Dryosaurus

Camptosaurus

Hypsilophodontidae

Thescelosaurus

Yandusaurus

Othnielia

Parksosaurus

Hypsilophodon

Zephyrosaurus

Orodromeus

The following cladogram of hypsilophodont relationships depicts the paraphyletic hypotheses; the "natural Hypsilophodontidae" hypothesis has been falling out of favor since the mid-late 1990s. It is after Brown et al. (2013), the most recent analysis of hypsilophodonts.[16] Ornithischia, Ornithopoda, and Iguanodontia were not designated in their result, and so are left out here. Additional ornithopods beyond Tenontosaurus are omitted. Dinosaurs traditionally described as hypsilophodonts are found from Agilisaurus or Hexinlusaurus to Hypsilophodon or Gasparinisaura.

Proctor Lake hypsilophodont
Skeleton of Convolosaurus (the Proctor Lake hypsilophodont)
unnamed

Heterodontosaurus

Scutellosaurus

Lesothosaurus

Agilisaurus

Hexinlusaurus

Othnielosaurus

Thescelosauridae
Orodrominae

TMP 2008.045.0002

Oryctodromeus

Albertadromeus

Orodromeus

Zephyrosaurus

Thescelosaurinae

Parksosaurus

Changchunsaurus

Haya

Jeholosaurus

Thescelosaurus assiniboiensis

Thescelosaurus neglectus

Hypsilophodontidae

Hypsilophodon

Gasparinisaura

Tenontosaurus

Other ornithopods

Norman's Hypsilophodontia

A more recent alternate phylogeny, by Norman in 2014, resolved a monophyletic Hypsilophodontia (the family Hypsilophodontidae was not used because of its history). Hypsilophodon grouped with Rhabdodontidae and Tenontosaurus.[15]

Clypeodonta
Hypsilophodontia

Hypsilophodon

Tenontosaurus

Rhabdodontidae

Iguanodontia

Dryosauridae

Ankylopollexia

Camptosaurus

Styracosterna

References

  1. ^ Dollo, L. (1982). "Première note sur les dinosauriens de Bernissart". Bulletin du Musée d'Histoire Naturelle de Belgique. 1: 161–180.
  2. ^ Sues, Hans-Dieter; Norman, David B. (1990). "Hypsilophodontidae, Tenontosaurus, Dryosauridae". In Weishampel, David B.; Dodson, Peter; Osmólska Halszka (eds.). The Dinosauria (1st ed.). Berkeley: University of California Press. pp. 498–509. ISBN 978-0-520-06727-1.
  3. ^ a b c Weishampel, David B.; Heinrich, Ronald E. (1992). "Systematics of Hypsilophodontidae and Basal Iguanodontia (Dinosauria: Ornithopoda)". Historical Biology. 6 (3): 159–184. doi:10.1080/10292389209380426.
  4. ^ Scheetz, Rodney D. (1998). "Phylogeny of basal ornithopod dinosaurs and the dissolution of the Hypsilophodontidae". Journal of Vertebrate Paleontology. 18 (3, Suppl): 1–94. doi:10.1080/02724634.1998.10011116.
  5. ^ Winkler, Dale A.; Murry, Phillip A.; Jacobs, Louis L. (1998). "The new ornithopod dinosaur from Proctor Lake, Texas, and the deconstruction of the family Hypsilophodontidae". Journal of Vertebrate Paleontology. 18 (3, Suppl): 87A. doi:10.1080/02724634.1998.10011116.
  6. ^ Buchholz, Peter W. (2002). "Phylogeny and biogeography of basal Ornithischia". The Mesozoic in Wyoming, Tate 2002. Casper, Wyoming: The Geological Museum, Casper College. pp. 18–34.
  7. ^ Weishampel, David B.; Jianu, Coralia-Maria; Csiki, Z.; Norman, David B. (2003). "Osteology and phylogeny of Zalmoxes (n.g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania". Journal of Systematic Palaeontology. 1 (2): 1–56. doi:10.1017/S1477201903001032.
  8. ^ Norman, David B.; Sues, Hans-Dieter; Witmer, Larry M.; Coria, Rodolfo A. (2004). "Basal Ornithopoda". In Weishampel, David B.; Dodson, Peter; Osmólska Halszka (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 393–412. ISBN 978-0-520-24209-8.
  9. ^ Varricchio, David J.; Martin, Anthony J.; Katsura, Yoshihiro (2007). "First trace and body fossil evidence of a burrowing, denning dinosaur" (PDF). Proceedings of the Royal Society B: Biological Sciences. 274 (1616): 1361–1368. doi:10.1098/rspb.2006.0443. PMC 2176205. PMID 17374596. Retrieved 2007-03-22.
  10. ^ Boyd, Clint A.; Brown, Caleb M.; Scheetz, Rodney D.; Clarke, Julia A. (2009). "Taxonomic revision of the basal neornithischian taxa Thescelosaurus and Bugenasaura". Journal of Vertebrate Paleontology. 29 (3): 758–770. doi:10.1671/039.029.0328.
  11. ^ a b Boyd, Clint A. (2015). "The systematic relationships and biogeographic history of ornithischian dinosaurs". PeerJ. 3 (e1523): e1523. doi:10.7717/peerj.1523. PMC 4690359. PMID 26713260. Retrieved 5 January 2016.
  12. ^ Gasulla, José Miguel; Escaso, Fernando; Narváez, Iván; Ortega, Francisco; Sanz, José Luis (2015). "A New Sail-Backed Styracosternan (Dinosauria: Ornithopoda) from the Early Cretaceous of Morella, Spain". PLoS ONE. 10 (12): e0144167. doi:10.1371/journal.pone.0144167. PMC 4691198. PMID 26673161.
  13. ^ Butler, Richard J.; Smith, Roger M.H.; Norman, David B. (2007). "A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia". Proceedings of the Royal Society B: Biological Sciences. 274 (1621): 2041–6. doi:10.1098/rspb.2007.0367. PMC 2275175. PMID 17567562.
  14. ^ Butler, Richard J.; Upchurch, Paul; Norman, David B. (2008). "The phylogeny of the ornithischian dinosaurs". Journal of Systematic Palaeontology. 6 (1): 1–40. doi:10.1017/S1477201907002271.
  15. ^ a b Norman, D.B. (2014). "On the history, osteology, and systematic position of the Wealden (Hastings group) dinosaur Hypselospinus fittoni (Iguanodontia: Styracosterna)". Zoological Journal of the Linnean Society. 173: 92–189. doi:10.1111/zoj.12193.
  16. ^ a b Brown, C. M.; Evans, D. C.; Ryan, M. J.; Russell, A. P. (2013). "New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta". Journal of Vertebrate Paleontology. 33 (3): 495–520. doi:10.1080/02724634.2013.746229.
  17. ^ a b Gilmore, C.W. (1915). "Osteology of Thescelosaurus, an orthopodous dinosaur from the Lance Formation of Wyoming". Proceedings of the United States National Museum. 49 (2127): 591–616. doi:10.5479/si.00963801.49-2127.591.
  18. ^ Zittel, K.A. von (1911). Grundzüge der Paläontologie (Paläzoologie) II. Abtielung Vertebrata (in German) (2 ed.). Berlin and München: Druck und verlad von R. Oldenbourg. p. 289.
  19. ^ Swinton, W.E. (1936). "Notes on the Osteology of Hypsilophodon, and on the Family Hypsilophodontidae". Journal of Zoology. 106 (2): 555–578. doi:10.1111/j.1469-7998.1936.tb08518.x.
  20. ^ Sternberg, C.M. (1940). "Thescelosaurus edmontonensis, n. sp., and classification of the Hypsilophodontidae". Journal of Paleontology. 14 (5): 481–494. JSTOR 1298552.
  21. ^ Galton, P.M. (1972). "Classification and Evolution of Ornithopod Dinosaurs". Nature. 239 (5373): 464–466. doi:10.1038/239464a0.
Albian

The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous epoch/series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 Ma (million years ago). The Albian is preceded by the Aptian and followed by the Cenomanian.

Aralosaurini

Aralosaurini is a tribe of basal lambeosaurine hadrosaurs endemic to Eurasia. It currently contains Aralosaurus (from the Aral sea of Kazakhstan) and Canardia (from Toulouse, Southern France).

Canardia

Canardia is an extinct genus of aralosaurin lambeosaurine dinosaur known from the Late Cretaceous Marnes d’Auzas Formation (late Maastrichtian stage) of Toulouse, Haute-Garonne Department, southern France. The type species Canardia garonnensis was first described and named by Albert Prieto-Márquez, Fabio M. Dalla Vecchia, Rodrigo Gaete and Àngel Galobart in 2013.

Convolosaurus

Convolosaurus (meaning "flocking lizard" after the concentration of juvenile fossils found) is a genus of basal ornithopod dinosaur from the Twin Mountains Formation from Proctor Lake in Comanche County, Texas. The type and only species is Convolosaurus marri.

Dryosaurus

Dryosaurus ( DRY-o-SAWR-əs; meaning 'tree lizard', Greek δρυς/drys meaning 'tree, oak' and σαυρος/sauros meaning 'lizard'; the name reflects the forested habitat, not a vague oak-leaf shape of its cheek teeth as is sometimes assumed) is a genus of an ornithopod dinosaur that lived in the Late Jurassic period. It was an iguanodont (formerly classified as a hypsilophodont). Fossils have been found in the western United States (and supposedly the Marnes de Bleville locality in Europe), and were first discovered in the late 19th century. Valdosaurus canaliculatus and Dysalotosaurus lettowvorbecki were both formerly considered to represent species of Dryosaurus.

Elasmaria

Elasmaria is a clade of iguanodont ornithopods known from Cretaceous deposits in South America, Antarctica, and Australia.

Fulgurotherium

Fulgurotherium (meaning "Lightning Beast") is the name given to a genus of dinosaur from the Early Cretaceous (Albian) Griman Creek Formation. It lived in what is now Australia.

The type species, Fulgurotherium australe, was named by Friedrich von Huene in 1932. The genus name is derived from Latin fulgur, "lightning", and Greek therion, "beast", a reference to the Lightning Ridge site in New South Wales. The specific name means "southern" in Latin. The holotype is BMNH R.3719, the opalised lower end of a femur, indicating a total body length of 1 to 1.5 metres.

Von Huene thought it was a theropod, a member of the Ornithomimidae. It has later been described as a hypsilophodont, a primitive ornithopod. However, this was based on a contentious reference of bones found in the Dinosaur Cave, leading to a possible confusion between multiple species of Euornithopoda. Most researchers today consider it a nomen dubium.

Its name is an unusual example of a name in which -therium was used for an animal which is not an extinct mammal.

Jaxartosaurus

Jaxartosaurus is a genus of hadrosaurid dinosaur similar to Corythosaurus which lived during the Late Cretaceous. Its fossils were found in Kazakhstan.

Lapampasaurus

Lapampasaurus is an extinct genus of hadrosaurid known from the Late Cretaceous Allen Formation (late Campanian or early Maastrichtian stage) of La Pampa Province, Argentina. It contains a single species, Lapampasaurus cholinoi.The generic name refers to the Argentine province of La Pampa. The specific name honours the late collector José Cholino. The material includes cervical, dorsal, sacral and caudal vertebrae, the forelimb girdle, and the partial hindlimb.

Loncosaurus

Loncosaurus (meaning uncertain; either Araucanian "chief" or Greek "lance" "lizard") was a genus of ornithopod dinosaur from the Upper Cretaceous of Provincia de Santa Cruz, Argentina. The type (and only known) species is Loncosaurus argentinus, described by the famous Argentinian paleontologist Florentino Ameghino, but is considered a dubious name. Details on this animal are often contradictory, befitting a genus that was long confused for a theropod.

Notohypsilophodon

Notohypsilophodon (meaning "southern Hypsilophodon") is a genus of euornithopod dinosaur from the Late Cretaceous of Argentina. It was described as the only "hypsilophodont" known from South America, although this assessment is not universally supported, and Gasparinisaura is now believed to have been a basal euornithopod as well.

Oryctodromeus

Oryctodromeus (meaning "digging runner") was a genus of small parksosaurid dinosaur. Fossils are known from the middle Cretaceous Blackleaf Formation of southwestern Montana and the Wayan Formation of southeastern Idaho, USA, both of the Cenomanian stage, roughly 95 million years ago. A member of the small, presumably fast-running herbivorous family Parksosauridae, Oryctodromeus is the first dinosaur published that shows evidence of burrowing behavior.

Othnielosaurus

Othnielosaurus is a genus of ornithischian dinosaur that lived about 155 to 148 million years ago, during the Late Jurassic-age Morrison Formation of the western United States. It is named in honor of famed paleontologist Othniel Charles Marsh, and was formerly assigned to the genus Laosaurus. This genus was coined to hold fossils formerly included in Othnielia, which is based on remains that may be too sparse to hold a name. O.C. Marsh named several species and genera in the late 19th century that have come to be recognized as hypsilophodonts or hypsilophodont-like animals, including Nanosaurus agilis, "N." rex (Othnielia), Laosaurus celer, L. consors, and L. gracilis. This taxonomy has become very complicated, with numerous attempts at revision in the years since; Othnielosaurus is part of decades of research to untangle the taxonomy left behind by Marsh and his rival Edward Drinker Cope from the Bone Wars. Othnielosaurus has usually been classified as a hypsilophodont, a type of generalized small bipedal herbivore or omnivore, although recent research has called this and the existence of a distinct group of hypsilophodonts into question.

Palo Cedro, California

Palo Cedro (Cedarwood) is a census-designated place (CDP) in Shasta County, California, United States. The population was 1,269 at the 2010 census, up from 1,247 at the 2000 census. It is 8 miles (13 km) east of Redding, California. The communities of Bella Vista (pop. 2,781), Millville (pop. 727), Shingletown (pop. 2,283), Oak Run (pop. 880), Whitmore (pop. 999), and Round Mountain (pop. 155) lie within a 15-mile (24 km) radius.

Originally, indigenous Native Americans lived in Northern California, including what is now Shasta County, prior to European American settlement. European American exploration of inland California started in 1769 and continued on into the 19th Century. Cow Creek, a Sacramento River tributary, that runs south through Palo Cedro, was a conduit for entrance into the Sacramento Valley by Hudson Bay Fur Company trappers including Alexander McLeod (1829) and John Work (1832). The town is named after cedarwood trees originally indigenous to the area in the 19th Century.

As of the 2010 census, Palo Cedro has a population density of 338.1 people per square mile (130.5/km2). Award winning country musician Merle Haggard lived in Palo Cedro for decades until his death on April 6, 2016.

Parksosaurus

Parksosaurus (meaning "William Parks's lizard") is a genus of hypsilophodont ornithopod dinosaur from the early Maastrichtian-age Upper Cretaceous Horseshoe Canyon Formation of Alberta, Canada. It is based on most of a partially articulated skeleton and partial skull, showing it to have been a small, bipedal, herbivorous dinosaur. It is one of the few described non-hadrosaurid ornithopods from the end of the Cretaceous in North America, existing around 70 million years ago.

Plesiohadros

Plesiohadros is an extinct genus of hadrosauroid dinosaur. It is known from a partial skeleton including the skull collected at Alag Teg locality, from the Campanian Djadochta Formation of southern Mongolia. The type species is Plesiohadros djadokhtaensis.

Rhabdodon

Rhabdodon (meaning "fluted tooth") is a genus of ornithopod dinosaur that lived in Europe approximately 70–66 million years ago in the Late Cretaceous. It is similar in build to a very robust "hypsilophodont" (non-iguanodont ornithopod), though all modern phylogenetic analyses find this ("Hypsilophodontia/tidae") to be an unnatural grouping, and Rhabdodon to be a basal member of Iguanodontia.

South Polar region of the Cretaceous

The South Polar region of the Cretaceous comprised the continent of East Gondwana–modern day Australia and Antarctica–a product of the break-up of Gondwana. The southern region, during this time, was much warmer than it is today, ranging from perhaps 4–8 °C (39–46 °F) in the latest Cretaceous Maastrichtian in what is now southeastern Australia. This prevented permanent ice sheets from developing and fostering polar forests, which were largely dominated by conifers, cycads, and ferns, and relied on a temperate climate and heavy rainfall. Major fossil-bearing geological formations that record this area are: the Santa Marta and Sobral Formations of Seymour Island off the Antarctic Peninsula; the Snow Hill Island, Lopez de Bertodano, and the Hidden Lake Formations on James Ross Island also off the Antarctic Peninsula; and the Eumeralla and Wonthaggi Formations in Australia.

The South Polar region housed many endemic species, including several relict forms that had gone extinct elsewhere by the Cretaceous. Of the dinosaur assemblage, the most diverse were the small hypsilophodont-like dinosaurs. The South Polar region also was home to the last labyrinthodont amphibian, Koolasuchus. The isolation of Antarctica produced a distinct ecosystem of marine life called the Weddellian Province.

Thescelosaurus

Thescelosaurus ( THESS-il-ə-SOR-əs; ancient Greek θέσκελος-/theskelos- meaning "godlike", "marvelous", or "wondrous" and σαυρος/sauros "lizard") was a genus of small ornithopod dinosaur that appeared at the very end of the Late Cretaceous period in North America. It was a member of the last dinosaurian fauna before the Cretaceous–Paleogene extinction event around 66 million years ago. The preservation and completeness of many of its specimens indicate that it may have preferred to live near streams.

This bipedal ornithopod is known from several partial skeletons and skulls that indicate it grew to between 2.5 and 4.0 meters (8.2 to 13.1 ft) in length on average. It had sturdy hind limbs, small wide hands, and a head with an elongate pointed snout. The form of the teeth and jaws suggest a primarily herbivorous animal. This genus of dinosaur is regarded as a specialized basal ornithopod, traditionally described as a hypsilophodont, but more recently recognized as distinct from Hypsilophodon. Several species have been suggested for this genus. Three currently are recognized as valid: the type species T. neglectus, T. garbanii and T. assiniboiensis.

The genus attracted media attention in 2000, when a specimen unearthed in 1993 in South Dakota, United States, was interpreted as including a fossilized heart. There was much discussion over whether the remains were of a heart. Many scientists now doubt the identification of the object and the implications of such an identification.

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